Profundiconus Limpalaeri Sp. Nov. (Gastropoda, Conilithidae)

A new deep water species from the Philippines: Profundiconus limpalaeri sp. nov. (Gastropoda, Conilithidae)

Manuel J. TENORIO INBIO - Dept. CMIM y Química Inorgánica, Facultad de Ciencias, Torre Norte, 1ª Planta, Universidad de Cádiz, 11510 Puerto Real, Cádiz, SPAIN e-mail: [email protected]

Eric MONNIER Conservatoire National des Arts et Métiers, Département Chimie, Alimentation, Santé, Environnement et Risque 292, rue Saint-Martin, 75003 Paris, FRANCE. e-mail: [email protected]

ABSTRACT We describe shell morphology using the terminology Profundiconus limpalaeri sp. nov. is described from deep water established in Röckel, Korn & Kohn (1995). For morphometric material taken around Balut Island, Philippines. The new species comparisons, adult shells selected among available specimens had been previously misidentified as Profundiconus cakobaui in several private collections, as well as in the collections of (Moolenbeek, Röckel & Bouchet, 2008) or Kurodaconus darkini the MNHN, were measured with a digital caliper, and the (Röckel, Korn & Richard, 1993). The observed similarities and measurements rounded to 0.1 millimeter. All measurements differences of these taxa with the new species are discussed. are in a spreadsheet, deposited as supporting information at https://dl.dropboxusercontent.com/u/30741402/

BIBLIOGRAPHY/XT12/limpalaeri.xls. For comparison of shell INTRODUCTION morphometry, we have performed analysis of the covariance The genus Profundiconus Kuroda, 1956 has been (ANCOVA) for different shell parameters namely maximum recently reviewed in Tenorio & Castelin (2016), and currently diameter (MD), height of the maximum diameter (HMD) and comprises 27 species living in deep water. The species in this spire height (SH), using species hypotheses as factor, and shell genus are spread over the Indo-Pacific region, except for length (SL) as covariate. Statistical tests were carried out using Profundiconus emersoni (Hanna, 1963), which occurs in the STATGRAPHICS 5.1, once all the measurement sets passed the East Pacific region. However, the inclusion of this species in normality tests. Profundiconus is only provisional (Tenorio et al. 2012). Due to RADULAR PREPARATIONS their deep-sea habitat, these species are difficult to sample, We use the terminology for radular morphology of and most of them are rare in collections with few exceptions. Tenorio & Castelin (2016). Specimens of shells containing the However, species of Profundiconus are relatively common dried animal inside were digested in concentrated aqueous in material dredged by research vessels in deep waters. To a potassium hydroxide for 24 hours. The contents were flushed lesser extent, small and medium-sized shells of still unnamed out of the shell by injecting distilled water through the aperture species of Profundiconus also surface occasionally in nets of the shell by means of a syringe with an incurved needle. of fishermen working in deep-water. That is the case for a The resulting mixture was then placed in a Petri dish and species caught by tangle nets at depths of 100-450 meters in examined with the binocular microscope. The entire radula was the waters around Balut Island and the neighboring Sarangani removed with fine tweezers and rinsed with distilled water, then Island in Southern Mindanao, Philippines. This species has mounted on a slide using Aquatex (Merck) Mounting Medium, been known for some time, and has been circulating around and examined under the compound microscope. Photos were among dealers and collectors with the names “Conus darkini” or obtained with a Charge Coupled Device (CCD) camera attached “Profundiconus cf. cakobaui” (Raybaudi Massilia in Poppe 2008; to the microscope. Tenorio & Castelin 2016). We have now been able to gather a sufficiently large homogeneous sample of this new species to Abbreviations describe it. We have compared these specimens qualitatively Museums and institutions and quantitatively withProfundiconus cakobaui (Moolenbeek, MNHN: Muséum National d’Histoire Naturelle, Paris, France. Röckel & Bouchet, 2008), and also with Kurodaconus darkini HMNS: Houston Museum of Natural Science, Houston, USA. (Röckel, Korn & Richard, 1993). Although we did not have access EM: Eric Monnier reference collection, Paris, France. to preserved material in our lots, we have concluded that the LL: Loïc Limpalaër reference collection, Haudivillers, France. specimens under study obtained from Balut Island are not CR: Christophe Roux reference collection, Vitry sur Seine, conspecific with eitherP. cakobaui nor K. darkini, and represent France. in fact a distinct, new species which is described hereby with PK: Paul Kersten reference collection, Hoornaar, The the name Profundiconus limpalaeri sp. nov. Netherlands. MATERIAL AND METHODS WJF: William J. Fenzan reference collection, Norfolk, Virginia, The taxonomy used in the present article follows USA. Tucker & Tenorio (2009) with the updates and modifications Other included in Tucker & Tenorio (2013). Specimens were collected m: meter by local fishermen either by tangle nets, trawling or dredging, mm: millimeter in depths ranging from 100 m to 500 meters.

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Shell morphometry Radular morphometry Distribution and habitat: To date, specimens have only been found at the type locality (Balut Island) and the neighboring S maximum shell length S /T shell length/radular tooth L L L Sarangani Island, in 100 to 450 m depth. Most specimens length found correspond to empty shells. A few specimens have been collected alive, although the soft parts were not preserved for RD relative diameter TL/APL radular tooth size/anterior portion length study. RSH relative spire height 100B /AP 100 x blade length/ Etymology: The species is named after Loïc Limpalaër, a L L French seashell collector, cone expert and a good friend, who anterior portion length co-described with the second author about 20 new species PMD relative position of the of cones. maximum diameter Description of the shell: Morphometric parameters: L = 32–40 mm; RD = 0.56–0.65; RSH = 0.21–0.27; PMD = 0.81–0.88. Shell RESULTS shape is ventricosely conical to conical. This is a moderately small to medium sized species. The spire is moderate to high, SYSTEMATICS concave and slightly sigmoid in profile. The whorl tops are flat Family Conilithidae Tucker & Tenorio, 2009 in cross section. The paucispiral protoconch of 1.5 whorls is Genus Profundiconus Kuroda, 1956 white to creamy-yellow, porcellaneous and translucent, with a Profundiconus limpalaeri sp. nov. (Plate 1, Figs. 1–4; Plate 2, maximum diameter of 0.98 mm (Fig. 1). The early teleoconch Figs. 1–4; Plate 3, Figs. 3–6) whorls are white or tan, with small tubercles which disappear Conus darkini – Poppe 2008: pl. 616, fig. 2, 3 & 5 (nonC. darkini by whorl five. Early teleoconch whorls have three to four cords. Röckel, Korn & Richard, 1993). The cords increase in number to six, seven or even more in late teleoconch whorls, but fade slightly in the outer whorls near Profundiconus cf. cakobaui– Tenorio & Castelin 2016: p. 23, fig. the shoulder. The shoulder is angulated, with a weak ridge 9E (non C. cakobaui Moolenbeek, Röckel & Bouchet, 2008). present. The sides of the shell are very slightly convex below Type material: Holotype and 7 paratypes. See table for details. the shoulder and then straight, becoming very slightly concave near the anterior end. The last whorl is smooth except for 8-9 Type Collection Dimensions Locality spiral grooves around the basal quarter. The posterior notch is (mm) shallow and C-shaped. There is no anterior notch. Holotype MNHN- 35.8 x 16.4 Balut Island, Southern IM-2000- Mindanao, Philip- 32434 pines, 300 m, trawled. Paratype 1 EM 34.9 x 15.8 Balut Island, Southern Mindanao, Philip- pines, 100-150 m. Paratype 2 CR 33.8 x 16.1 Balut Island, Southern Mindanao, Philip- pines, 200-300 m, dredged. Paratype 3 PK 38.2 x 16.7 Sarangani Island, Southern Mindanao, Philippines, 300 m. Paratype 4 HMNS 34.0 x 16.5 Balut Island, Southern Mindanao, Philippines, 200-250 m. Paratype 5 WJF 32.0 x 15.0 Tinina, Balut Island, Southern Mindanao, Philippines, 500 m, by tangle net. Paratype 6 WFJ 35.0 x 16.4 Balut Island, Southern Mindanao, Philip- pines. Paratype 7 LL 36.0 x 16.9 Balut Island, Southern Mindanao, Philip- pines, 400 m.

Material examined: More than 10 additional specimens, mostly dead collected, have been studied; all of them from Balut Island, Southern Mindanao, Philippines, collected in deep water. These specimens are in private collections, most of them ex-collection Conchology Inc. Fig. 1.- Protoconch and early teleoconch whorls of P. limpalaeri

Type locality: Balut Island, Southern Mindanao, Philippines. sp. nov. (Holotype, SL 35.8 mm).

42  XENOPHORA TAXONOMY 12 - JUL 2016 M. TENORIO & E. MONNIER - A new deep water species from the Philippines: Profundiconus limpalaeri sp. nov. Ground color white to very pale cream. Early teleoconch Living animal and radula: whorls often white, patternless. Alternating brown blotches Details about the living animal are unknown. starting about whorl 4-5, extending over the late spire whorls reaching the shoulder. Last whorl overlaid with large irregular axial brown blotches and streaks in variable amount. The DISCUSSION pattern can be dense, covering most of the surface of the last Profundiconus limpalaeri sp. nov. is most similar to whorl (i.e. Plate 2, Fig 4a; Plate 3, Fig. 6). In some specimens, K. darkini (Plate 3, Fig. 2) and to P. cakobaui (Plate 3, Fig. 1). the axial brown blotches are arranged forming also narrow K. darkini can be easily separated from P. limpalaeri sp. nov. interrupted spiral bars (i.e. Plate 1, Fig. 4a; Plate 2, Fig 2; Plate by its larger adult size (51 – 87 mm versus 32 – 40 mm for P. 3, Fig. 4). The columella is white. The aperture is bluish-white, limpalaeri sp. nov.), larger number of strongly tuberculated usually stained with purplish-brown at the inner edge of the spire whorls (9 – 10 versus 5 in P. limpalaeri sp. nov.) and its lip, especially towards the base. The aperture may be slightly multispiral protoconch of 3.5 whorls (paucispiral of 1.5 whorls transparent allowing the outer pattern of the shell to add color in P. limpalaeri sp. nov.). It is important to point out that to the bluish-white aperture. The periostracum is yellowish, thin despite the external appearance of the shell, darkini is not a and transparent. The operculum (Fig. 2) received with paratype Profundiconus. A study of the radular morphology (Fig. 3A) 5 is oval-shaped, yellowish and translucent. Its relative length indicated that it is a typical Conidae sensu Tucker & Tenorio corresponds approximately to 17% of the aperture height. It (2009). The radular tooth is small: its total length relative to lacks the characteristic serrations on the outer margin observed shell length SL/TL = 89–90. The anterior portion is much shorter in the operculum of other species in genus Profundiconus than the posterior section of the tooth (TL/APL= 2.9). There is (Tenorio & Castelin 2016). Instead, it displays just rather gentle one barb and a rounded blade, covering approximately two- undulations on the margin. There is no absolute certainty, thirds (66%) of the anterior portion of the tooth. Serration with though, about whether or not this operculum actually belongs 18 to 23 small denticles arranged in two rows, ending in a small, to the specimen corresponding to paratype 5. rounded terminating cusp. Shaft fold absent. Basal spur present on top of the rounded base of the tooth. Recent molecular studies suggest that this species belongs to the same clade as Kurodaconus luciae (Puillandre, pers. comm.), and hence the attribution of darkini to genus Kurodaconus.

We are provisionally considering limpalaeri as a member of genus Profundiconus, pending confirmation by examination of radular morphology and/or DNA sequencing. On the other hand, we have been able to extract and examine the radular teeth of two specimens of P. cakobaui, confirming in Fig. 2.- Operculum of P. limpalaeri sp. nov. received with this way the attribution of this species to genusProfundiconus paratype 5. (Fig 3B, C). The radular tooth is rather large: its total length

relative to shell length SL/TL = 32–38. The anterior portion is

Fig. 3.- Radular teeth of: (A) Kurodaconus darkini (MNHN IM-2009-31324, Récif Pétrie, Nouvelle Calédonie, 448-464 m,

SL 57.3 mm); (B) Profundiconus cakobaui, paratype (MNHN IM-2000-21033, Ride de Lau, Somo-somo Strait, Fiji, 414-510 m, SL

18.4 mm); (C) Profundiconus cakobaui (MNHN IM-2008-1243, Bligh Water, Fiji, 567-699 m, SL 24.8 mm).

XENOPHORA TAXONOMY 12 - JUL 2016  43 M. TENORIO & E. MONNIER - A new deep water species from the Philippines: Profundiconus limpalaeri sp. nov.

  Fig. 4.- Plot of MD versus SL for P. limpalaeri sp. nov. ( ) and P. cakobaui ( ).

much shorter than the posterior section of the tooth (TL/APL= 3.2–3.8). There is one barb and a pointed, well-defined blade, which covers 40 to 60% of the anterior portion of the tooth. There is an external cusp located at approximately the lower quarter of the anterior portion of the tooth, extending between 80% and 95% of the length of the anterior portion of the tooth. The external cusp is laterally expanded and serrated, with 6 denticles. Immediately below the waist, there is a characteristic fringe composed of closely spaced projections pointing towards the apex. Shaft fold is present. A large and prominent basal spur is present on top of the slanted base of the tooth. All these features of the radular tooth of P. cakobaui are fully consistent with those reported for other species in genus Profundiconus (Tenorio & Castelin 2016). P. limpalaeri sp. nov. and P. cakobaui, known from Fiji and Tonga, exhibit an apparently similar shell shape, but

they differ clearly in mean shell length. Comparison of mean SL between the two species yields statistically significant results as

expected: mean SL for P. limpalaeri sp. nov. 36.1 mm; mean SL for P. cakobaui 17.4 mm (t = -13.5, p = 3.93 x 10-12; U = 128, p =

0.0001). ANCOVA (using SL as covariate) indicates that there are no significant differences between these two species in SH (F = 2.98, p = 0.099) or HMD (F = 2.07, p = 0.165), but they differ in MD (F = 8.65, p = 0.0078, Fig. 4).

Hence, P. limpalaeri sp. nov. has a significantly larger and broader shell in comparison with P. cakobaui. Apart from these differences in size and shape, the pattern and general features of the shell of the two species are similar, including the paucispiral protoconch with tuberculated early teleoconch whorls. However, the cords on the teleoconch whorls of P. Fig. 5.- Comparative views at the same scale of the cakobaui are less numerous than in P. limpalaeri sp. nov. (Fig. teleoconch whorls of: A) P. cakobaui (Paratype, MNHN IM- 5). In P. cakobaui, the subsutural area is essentially smooth, and 2000-21033, SL 18.4 mm); B) P. limpalaeri sp. nov. (Holotype)

44  XENOPHORA TAXONOMY 12 - JUL 2016 M. TENORIO & E. MONNIER - A new deep water species from the Philippines: Profundiconus limpalaeri sp. nov. then showing 3-4 cords (Fig. 5A). In P. limpalaeri sp. nov., there BIBLIOGRAPHY are more cords, and these are more evenly distributed (Fig. 5B). Kohn, A. J., Nishi, M. & Pernet, B. 1999. Snail spears and scimitars: a character analysis of Conus radular teeth. Journal Apart from these morphological features, P. limpalaeri of Molluscan Studies, 65: 461–481. sp. nov. and P. cakobaui are separated geographically by a Kuroda, T. 1956. New species of the Conidae (Gastropoda). distance of approximately 6300 kilometers. Since a paucispiral Venus, 19(1): 1–16. protoconch indicates a direct, non-planktonic mode of Moolenbeek, R. G., Röckel, D. & Bouchet, P. 2008. New records development, and the two taxa are reproductively isolated, and new species of cones from deeper water off Fiji (Mollusca, there is support for the hypothesis of two separate species. Gastropoda, Conidae). Vita Malacologica 6: 35–49. Raybaudi-Massilia, G. in Poppe, G. T. 2008. Philippine Marine ACKNOWLEDGMENTS Mollusks, Vol. II. Conchbooks, Hackenheim, Germany. Our most sincere thanks to: Prof. Philippe Bouchet, Virginie Röckel, D., Korn, W. & Kohn, A. J. 1995. Manual of the Living Heros and Philippe Maestrati from the MNHN, Paris, for all Conidae, Volume 1: Indo-Pacific Region: 1-517. Hackenheim. the facilities provided for the study of deep water cones from Verlag Christa Hemmen. Fiji and New Caledonia; Manuel Caballer, project E-RECOLNAT: Röckel, D., Korn, W., & Richard, G., 1993. From the Western ANR-11-INBS-0004, MNHN for the photos of the types of Pacific Conus darkini n. sp. (Prosobranchia: Conidae). La P. cakobaui; Guido T. Poppe and Conchology, Inc. for kindly Conchiglia: International Shell Magazine, 25 (267): 48–49. providing photos and information about specimens of the new Röckel, D., Richard, G. & Moolenbeek, R. G. 1995. Deep- species; special thanks to William J. Fenzan for loaning material water Cones (Gastropoda: Conidae) from the New Caledonia from his reference collection for study, and for suggesting region. Mémoires du Muséum National d’Histoire Naturelle, improvements to the manuscript. 167: 557–594. Tenorio, M. J. & Castelin, M. 2016. Genus Profundiconus Kuroda, 1956 (Gastropoda, Conoidea): Morphological and molecular studies, with the description of five new species from the Solomon Islands and New Caledonia. European Journal of Taxonomy 173:1–45. Tenorio M. J., Tucker J. K. & Chaney H. W. 2012. The Families Conilithidae and Conidae – The Cones of the Eastern Pacific.A Conchological Iconography, Vol. 18. Conchbooks, Hackenheim, Germany. Tucker, J. K. & Tenorio, M. J. 2009. Systematic Classification of Recent and Fossil Conoidean Gastropods, with Keys to the Genera of Cone Shells. ConchBooks, Hackenheim, Germany, 295 pp. Tucker, J. K. & Tenorio, M. J. 2013. Illustrated Catalog of the Living Cone Shells. MdM Publishing, Wellington FL, USA, 517 pp.

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Plate 1.- 1-4 Profundiconus limpalaeri sp. nov. Balut Is., Philippines. 1a-b. Paratype 1, dorsal and ventral views, EM coll., 34.9 x 15.8 mm; 2a-b. Holotype, dorsal and ventral views, MNHN IM-2000-32434, 35.8 x 16.4 mm. 3a-b. Paratype 2, dorsal and ventral views, CR coll., 33.8 x 16.1 mm. 4. Paratype 3, PK coll., 38.2 x 16.7 mm.

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Plate 2.- 1-4 Profundiconus limpalaeri sp. nov. Balut Is., Philippines. 1a-b. Paratype 4, dorsal and ventral views, HMNS, 34.0 x 16.5 mm; 2a-b. Paratype 5 (w/p, w/o), dorsal and ventral views, WJF coll., 32.0 x 15.0 mm. 3a-b. Paratype 6, dorsal and ventral views, WJF coll., 35.0 x 16.4 mm. 4. Private coll., ex-Conchology, Inc., 36.9 x 17.3 mm. XENOPHORA TAXONOMY 12 - JUL 2016  47 M. TENORIO & E. MONNIER - A new deep water species from the Philippines: Profundiconus limpalaeri sp. nov.

Plate 3.- 1a-b. Holotype of Conus cakobaui Moolenbeek, Röckel & Bouchet, 2008, MNHN IM-2000-21030, Somo-somo Strait, South of Vanua Levu, Fiji, 426-487 m, 18.9 x 8.5 mm; 2a-b. Holotype of Conus darkini Röckel, Korn & Richard, 1993, MNHN IM- 2000-2568, E Cap des Pins, Ride des Loyauté, Nouvelle Calédonie, 575 m, 55.6 x 24.4 mm. 3-6 Profundiconus limpalaeri sp. nov. Balut Is., Philippines, private coll., ex-Conchology, Inc. 3. 38.8 x 17.7 mm. 4. 38.1 x 18.5 mm. 5. 35.9 x 16.7 mm. 6. 37.5 x 17.4 mm.

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